Meat Science 66 (2003) 21–32

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Effects of fatty acids on meat quality: a review

J.D. Wood*, R.I. Richardson, G.R. Nute, A.V. Fisher,
M.M. Campo, E. Kasapidou, P.R. Sheard, M. Enser
Department of Clinical Veterinary Science, Division of Farm Animal Science,
University of Bristol, Langford, Bristol BS40 5DU, UK

Received 27 September 2002; accepted 11 November 2002

Abstract
Interest in meat fatty acid composition stems mainly from the need to find ways to produce healthier meat, i.e. with a higher ratio
of polyunsaturated (PUFA) to saturated fatty acids and a more favourable balance between n-6 and n-3 PUFA. In pigs, the drive
has been to increase n-3 PUFA in meat and this can be achieved by feeding sources such as linseed in the diet. Only when con-
centrations of a-linolenic acid (18:3) approach 3% of neutral lipids or phospholipids are there any adverse effects on meat quality,
defined in terms of shelf life (lipid and myoglobin oxidation) and flavour. Ruminant meats are a relatively good source of n-3
PUFA due to the presence of 18:3 in grass. Further increases can be achieved with animals fed grain-based diets by including whole
linseed or linseed oil, especially if this is ‘‘protected’’ from rumen biohydrogenation. Long-chain (C20–C22) n-3 PUFA are syn-
thesised from 18:3 in the animal although docosahexaenoic acid (DHA, 22:6) is not increased when diets are supplemented with
18:3. DHA can be increased by feeding sources such as fish oil although too-high levels cause adverse flavour and colour changes.
Grass-fed beef and lamb have naturally high levels of 18:3 and long chain n-3 PUFA. These impact on flavour to produce a ‘grass
fed’ taste in which other components of grass are also involved. Grazing also provides antioxidants including vitamin E which
maintain PUFA levels in meat and prevent quality deterioration during processing and display. In pork, beef and lamb the melting
point of lipid and the firmness/hardness of carcass fat is closely related to the concentration of stearic acid (18:0).
# 2003 Elsevier Ltd. All rights reserved.
Keywords: Cattle; Fatty acids; Meat quality; Pigs; Sheep

1. Introduction increased to above 0.4. Since some meats naturally have

There has been an increased interest in recent years in
ways to manipulate the fatty acid composition of meat.
This is because meat is seen to be a major source of fat
in the diet and especially of saturated fatty acids, which
have been implicated in diseases associated with modern
life, especially in developed countries. These include
various cancers and especially coronary heart disease. In
the UK, the Department of Health (1994) recommended
that fat intake be reduced to 30% of total energy intake
(from about 40%) with a figure of 10% of energy intake
for saturated fatty acids (from 15%). At the same time,
the recommended ratio of polyunsaturated fatty acids
(PUFA) to saturated fatty acids (P:S) should be
* Corresponding author. Tel.: +44-117-928-9293; fax: +44-117-
928-9324.
E-mail address: jeff.wood@bristol.ac.uk (J.D. Wood).
0309-1740/03/$ - see front matter # 2003 Elsevier Ltd. All rights reserved.
doi:10.1016/S0309-1740(03)00022-6
a P:S ratio of around 0.1, meat has been implicated in
causing the imbalanced fatty acid intake of today’s
consumers. For this reason, ways to improve the P:S
ratio during meat production are required. More
recently, nutritionists have focussed on the type of
PUFA and the balance in the diet between n-3 PUFA
formed from a-linolenic acid (18:3) and n-6 PUFA
formed from linoleic acid (18:2) (Williams, 2000). The
ratio of n-6:n-3 PUFA is also a risk factor in cancers
and coronary heart disease, especially the formation of
blood clots leading to a heart attack (Enser, 2001). The
recommendation is for a ratio of less than 4 and again
some meats are higher than this. As with the P:S ratio,
meats can be manipulated towards a more favourable
n-6:n-3 ratio.
The increasing awareness of the need for diets to
contain higher levels of n-3 PUFA has focused on the
importance of meat as a natural supplier of these to the
22 J.D. Wood et al. / Meat Science 66 (2003) 21–32

diet. The ratio of n-6:n-3 PUFA is particularly bene- Table 1

ficial (low) in ruminant meats, especially from animals Fat and fatty acid composition of beef, lamb and pork loin steaks
purchased from four supermarkets (Enser et al., 1996)
that have consumed grass which contains high levels of
18:3. Ruminants also naturally produce conjugated Beef Lamb Pork
linoleic acids (CLAs) which may have a range of nutri-
Whole steak
tional benefits in the diet (Enser, 2001). Fata 15.6 30.2 21.1
Fatty acids are involved in various ‘‘technological’’
aspects of meat quality. Because they have very different Muscleb
melting points, variation in fatty acid composition has 16:0 palmitic 25.0 22.2 23.2
an important effect on firmness or softness of the fat in 18:0 stearic 13.4 18.1 12.2
18:1 n-9 oleic 36.1 32.5 32.8
meat, especially the subcutaneous and intermuscular 18:2 n-6 linoleic 2.4 2.7 14.2
(carcass fats) but also the intramuscular (marbling) fat. 18:3 n-3 a-linolenic 0.70 1.37 0.95
Groups of fat cells containing solidified fat with a high 20:4 n-6 arachidonic 0.63 0.64 2.21
melting point appear whiter than when liquid fat with a 20:5 n-3 EPA 0.28 0.45 0.31
lower melting point is present, so fat colour is another 22:6 n-3 DHA 0.05 0.15 0.39
Total fatty acidsc 3.8 4.9 2.2
aspect of quality affected by fatty acids. The ability of P:S 0.11 0.15 0.58
unsaturated fatty acids, especially those with more than n-6:n-3 2.11 1.32 7.22
two double bonds, to rapidly oxidise, is important in
regulating the shelf life of meat (rancidity and colour Fatb
deterioration). However, this propensity to oxidise is 16:0 palmitic 26.1 21.9 23.9
18:0 stearic 12.2 22.6 12.8
important in flavour development during cooking. 18:1 n-9 oleic 35.3 28.7 35.8
The aim of this article is to summarise the main effects 18:2 n-6 linoleic 1.1 1.3 14.3
of fatty acid composition on meat quality and to review 18:3 n-3 a-linolenic 0.48 0.97 1.43
recent work showing the effects on meat quality of chan- C20–C22 n-3 PUFAd ND ND 0.36
ges in fatty acid composition achieved during production. ND, not detectable.
a
% Of steak.
b
% Of total fatty acids.
c
2. Fatty acid composition of meat % Of muscle.
d
% Of fat tissue.
A survey conducted by Enser, Hallett, Hewett, Fur-
sey, and Wood (1996) illustrated the differences in fatty Table 2
acid composition and content between beef, lamb and Concentrations of neutral lipids and phospholipids in longissimus
pork. Fifty loin steaks or chops from each species were muscle and % of 18:2 and 18:3 in each fraction in steers (Choi, Enser,
Wood, & Scollan, 2000) and gilts (Enser et al., 2000)
purchased from four supermarkets to represent the
meat on sale to the public (Table 1). The total fat con- Neutral Phospholipids PL/NL
tent of the steaks (obtained by dissection) was highest in lipids
lamb, probably because of a lower level of fat trimming Steers (Welsh Concentrationa 2.56 0.53
during butchery. The total fatty acid composition of the Black controls) 18:2 1.05 12.89 12.3
longissimus muscle, including some fat attached to the 18:3 0.48 2.89 6.0
perimysium, was also highest in lamb and was least in
Gilts (Large Concentrationa 0.69 0.40
pork. The most obvious difference in fatty acid compo-
White controls) 18:2 10.10 30.20 3.0
sition was that 18:2 was higher in pork, causing a higher 18:3 0.81 0.90 1.1
P:S ratio. This is due to the high content of 18:2 in the
a
cereal-based diets consumed by meat animals and this % Of muscle.
produced an undesirably high n-6:n-3 ratio. The rumi-
nant meats had a more favourable n-6:n-3 ratio, due
both to less18:2 than in pork and relatively high levels of control animals from two studies are compared. The
of n-3 PUFA, especially 18:3. The study also showed percentage of 18:2 in longissimus phospholipids was 12
that the long chain (C20–C22) n-3 PUFA were at low times greater than that in neutral lipids in the steers and
but significant levels in pork subcutaneous fat but not 3 times greater in pigs. Ratios were 6 and 1 for 18:3 in
detectable in beef and lamb, reflecting a relatively the steers and pigs, respectively.
greater deposition of long chain derivatives of 18:3 in Differences in muscle fibre type between muscles are
pig neutral lipids (triacylglycerols). In ruminant muscle reflected in differences in fatty acid composition. ‘‘Red’’
and adipose tissue, PUFA are restricted almost exclu- muscles have a higher proportion of phospholipids than
sively to the phospholipid fraction. An illustration of ‘‘white’’ muscles and therefore a higher percentage of
this species effect on 18:2 and 18:3 is in Table 2. Groups PUFA. An example is given in Table 3 comparing the
 J.D. Wood et al. / Meat Science 66 (2003) 21–32
Table 3
Fatty acid content and composition of total lipid in longissimus and
the ‘‘redder’’ gluteobiceps muscle in 30 grass-fed steers (Enser et al.,
1998)
Longissimus
Total fatty acidsa 2.9
18:0 15.8
18:1 n-9 34.7
18:2 n-6 2.5
18:3 n-3 1.2
20:4 n-6 0.8
20:5 n-3 0.5
22:6 n-3 0.07
P:S 0.08
n-6:n-3 1.32
a
% Of muscle.
longissimus muscle with the redder leg muscle gluteobi-
ceps. The muscles were taken from 30 grass-fed steers
examined by Enser, Hallett, Hewett, Fursey, Wood, and
Harrington (1998). The P:S ratio was significantly
higher in gluteobiceps due to higher concentrations of
most PUFA.
Studies on poultry meat have shown similarities with
pork i.e. the meat fatty acids are relatively unsaturated
although 18:2 is at a higher level (Enser, 1999). This
similarity with pork has caused the US pig industry to
label pork as the ‘‘other white meat’’.
3. Manipulation of fatty acid composition and the
effects on meat quality
3.1. Components of meat quality
The components of technological meat quality influ-
enced by fatty acids are fat tissue firmness (hardness),
shelf life (lipid and pigment oxidation) and flavour.
Although there have been suggestions that dietary fatty
acids influence tenderness and juiciness, these are more
likely to be affected by the total amount of fatty acids
rather than individual ones. The effect of fatty acids on
firmness is due to the different melting points of the
fatty acids in meat. In the 18C fatty acid series, stearic
acid (18:0) melts at 69.6  C, oleic acid (18:1) at 13.4  C,
18:2 at 5  C and 18:3 at 11  C. Thus as unsaturation
increases, melting point declines. Variation in the struc-
ture of the molecule are also important. For example,
trans fatty acids melt at a higher temperature than their
cis-isomers and branched chain fatty acids have lower
melting points than the straight chain fatty acids with
the same number of carbon atoms (Enser, 1984).
The effect of fatty acids on shelf life is explained by
the propensity of unsaturated fatty acids to oxidise,
leading to the development of rancidity as display times
increases. The colour change is due to the oxidation of
23
red oxymyoglobin to brown metmyoglobin, this reac-
tion generally proceeding in parallel to that of rancidity.
Several studies have shown that lipid oxidation pro-
ducts can promote pigment oxidation and vice versa
Gluteobiceps although the strength of the relationship between these
3.4
two aspects of shelf life is sometimes low (Renerre,
14.3 2000). Antioxidants, especially a-tocopherol (vitamin E)
35.7 have been used to delay lipid and colour oxidation and
3.3 to extend shelf life.
1.5 The effect of fatty acid on meat flavour is due to the
1.3
0.8
production of volatile, odourous, lipid oxidation pro-
0.13 ducts during cooking and the involvement of these with
0.13 Maillard reaction products to form other volatiles
1.35 which contribute to odour and flavour. The unsaturated
phospholipid fatty acids are particularly important in
flavour development. Early research showed that the fat
tissues in meat were the source of the characteristic
species flavour (Mottram, 1998).
3.2. Fatty acids and meat quality in pigs
Studies conducted at Bristol in the 1970s and 1980s
showed that 18:0 and 18:2 are particularly important
contributors to fat tissue firmness. As fatty acid com-
position was changed for reasons of diet, genetics, sex
or fatness, these two showed the highest correlations
with firmness measured subjectively (a finger test) or
objectively (using a compression tester or penet-
rometer). The ratio of 18:0:18:2 was found by Whit-
tington, Prescott, Wood, and Enser (1986) to provide
the best prediction of firmness. In a study by Wood,
Enser, MacFie, Smith, Chadwick, and Ellis (1978) of
pigs selected for lean content, the melting point of
extracted lipid was also closely related to the con-
centrations of 18:0 and 18:2, with 18:0 showing the
highest correlation. The role of 18:0 is particularly
interesting since it varies across a smaller range than
18:2.
Fat tissue consists not only of fatty acids contained in
fat cells but also the connective tissue matrix associated
with water. In pigs, the concentration of water in thin
(i.e. underdeveloped) backfat is very high as is that of
collagen (Table 4). These constituents are also good
predictors of tissue firmness along with the concentra-
tions of 18:0 and 18:2 and the thickness of the fat tissue
itself. These results show that fat tissue development in
the pig is an extremely ordered process with all the
major constituents closely interrelated.
As subcutaneous fat tissue develops and changes in
composition, it becomes more cohesive i.e. is less easily
separated within itself layer by layer. The problem of fat
tissue separation is unsightly in fresh pork and particu-
larly in bacon or ham. Studies have shown that cohe-
siveness is closely related to water, collagen, 18:0 and
18:2 fatty acid concentrations, just as is firmness
(Table 4).
24 J.D. Wood et al. / Meat Science 66 (2003) 21–32
Table 4
Composition and quality of backfat in entire and castrated male pigs
given different diets, and correlations between tissue constituents and
fat quality (Wood et al., 1985)
Lean entires Fat castrates
P2 fat thickness (mm) 9.0 15.4
Firmness (N) 1.0 3.4
Cohesiveness (N/m2103) 2.5 5.8
Water (mg/g) 257 151
18:0a 11.3 14.6
18:2a 29.9 9.3
Correlations between characteristics
P2 Water 18:0
Firmness +0.67 0.45 +0.70
Cohesiveness +0.47 0.48 +0.52
a
% Of total fatty acids.
After slaughter, various changes occurring in pig meat
favour oxidation, for example the release of iron from
within cells and the loss of the glutathione peroxidase
enzyme system (Morrissey, Sheehy, Galvin, Kerry, &
Buckley, 1998). Because the lipids in pig meat are rela-
tively unsaturated, attempts to further increase con-
centrations of PUFA risk the generation of lipid
oxidation products, leading to off-odours and flavours
and colour changes.
Several papers have examined the effects of dietary
oils containing a high proportion of 18:2 on the fatty
acid composition and quality of pigmeat. Examples of
such oils are soya, peanut, maize and sunflower. Con-
centrations of 18:2 can easily be raised from basal levels
of around 10–15% of fatty acids to over 30% (Hart-
man, Costello, Libal, & Walhlstrom, 1985; West &
Myer, 1987), the effect being rapidly achieved. For
example in one study, Warnants, Van Oeckel, and
Boucqué (1999) showed the full effect of feeding full fat
soya oil to pigs from 30 kg live weight was achieved in 6
weeks and 50% of this was achieved in 2 weeks.
Larick, Turner, Schoenherr, Coffey, and Pilkington
(1992) showed that muscles with raised levels of 18:2
oxidised rapidly when heated, producing various vola-
tile compounds, including the aldehydes pentanal and
hexanal. However, there was no change in the flavour of
ground pork patties made from meat with low or high
concentrations of 18:2 and no effect on muscle colour.
Hartman et al. (1985) raised the 18:2 concentration
from 15 to 33% of total fatty acids and also found no
effects on the flavour or colour of pork chops. A similar
result was found by West and Myer (1987). Melton
(1990) speculated that because pork was naturally high
in 18:2, its oxidation products are recognised by con-
sumers as natural components of pork flavour.
Supplementing pig diets with 18:3 to lower the n-6:n-3
ratio has been examined by several workers. Canola
(rapeseed) oil and especially flaxseed (linseed) are good
sources of this. In some papers, no effects on meat
quality parameters have been observed (Enser,
Richardson, Wood, Gill, & Sheard, 2000; Leskanich,
Matthews, Warkup, Noble, & Hazzledine, 1997) but in
others, adverse effects on odours and flavours have been
detected, especially when oxidative stress in meat is
increased by preparation treatment (e.g. salt injection,
comminution, freezing and reheating; Myer, Johnson,
Knauft, Gorbet, Brendemuhl, & Walker, 1992; Shack-
elford, Reagan, Haydon, & Miller, 1990). A possible
explanation for these results lies in the level of 18:3
achieved in the fat or muscle tissue. Above about 3% of
18:2
total fatty acids, the mix of volatile compounds pro-
0.60 duced on cooking seems to adversely impact on flavour
0.53 as detected by taste panellists. Campo, Nute, Wood,
Elmore, Mottram, and Enser (in press), in a model sys-
tem, found that odour scores were often higher for 18:3
than for 18:2, i.e. 18:3 produces more intense odours.
Enser et al. (2000) and Sheard, Enser, Wood, Nute,
Gill, and Richardson (2000) showed that the n-6:n-3
ratio in pork could be reduced close to the target level
(less than 4) by feeding crushed whole linseed, with no
detectable adverse effects on meat quality. They showed
that increasing the 18:3 content of longissimus muscle
total lipid from 1% in controls to 1.6% in those fed
linseed, which lowered the n-6:n-3 ratio to five (com-
pared with nine in controls), had no effects on lipid
oxidation measured using thiobarbituric acid reacting
substances test (TBARSs), colour saturation (vividness
of red colour) or the eating quality of grilled loin chops.
Kouba, Enser, Whittington, Nute, and Wood (in
press) showed that feeding a diet containing 6% whole
crushed linseed reduced the n-6:n-3 ratio in longissimus
muscle to 3.9 in only 20 days from 40 kg live weight
compared with a ratio of 7.6 in controls. Ratios were 3.0
at 60 days and 3.1 at 100 days on the linseed diet, cor-
responding to 18:3 concentrations of 3.0 and 2.2% in
longissimus total lipid at 60 and 100, days respectively.
This supports the conclusion of Warnants et al. (1999)
that maximal feeding effects with essential fatty acids
can be achieved within 40 days, with half of the effect
occurring within 2 weeks. Results in Fig. 1a from the
study of Kouba et al. show the similarity of 18:3 con-
centrations in pig neutral lipids and phospholipids,
whereas the long-chain PUFAs, e.g. eicosapentaenoic
acid (EPA, 20:5) are much higher in phospholipids. For
both fatty acids, the concentration at 100 days on the
linseed diet was significantly lower than at 60 days.
Levels of n-3 PUFA in the pigs fed the linseed diet pro-
duced higher TBARS levels following conditioning for
10 days followed by simulated retail display for a fur-
ther 7 days (Fig. 1b) but this did not impact on muscle
colour (saturation) during the display period (Fig. 1c).
However, there were detectable differences in odours
and flavours between the diets in grilled pork chops as
identified by the trained taste panel (Fig. 1d). The pigs
 J.D. Wood et al. / Meat Science 66 (2003) 21–32 25

Fig. 1. Effect of a 6% whole crushed linseed diet on fatty acid composition and meat quality in pigs (Kouba et al., in press). (a) 18:3 (top) and 20:5
(bottom) as % longissimus neutral lipids (NL) and phospholipids (PL). (b) Lipid oxidation in longissimus after 10 days conditioning at 1  C and 7
day display in over-wrapped packs at 4  C. Assessed as TBARS (mg malonaldehyde/kg). (c) Colour saturation in longissimus during retail display in
overwrapped packs. (d) Taste panel results for griddled loin steaks (1–8 scales).

fed linseed had lower scores for pork odour and pork
flavour and higher scores for abnormal odour and
abnormal flavour. These results confirm the US data
(e.g. Shackelford et al., 1990) showing that 18:3 con-
centrations of above about 3% in total lipid or neutral
lipid can produce relatively undesirable flavours if pro-
cessing conditions accelerate lipid oxidation.
In the study of Kouba et al. (in press; Fig. 1), the
concentration of vitamin E in longissimus was lower in
the pigs fed linseed after 60 days than in controls (2.1 vs.
2.9 mg/g). Vitamin E added to the diet was the same in
both groups (150 mg/g) so this effect could be associated
with greater utilisation of the antioxidant to protect the
more unsaturated phospholipid fatty acids.
Changes in fatty acid composition have not been
directly linked to changes in myoglobin oxidation and
muscle colour in many of the pig studies reported
(Kouba et al., in press; Larick et al., 1992; Sheard et al.,
2000; West & Myer, 1987). In several studies in which
supranutritional vitamin E has been used to inhibit fatty
acid oxidation, concomitant improvements in colour
stability have also not been observed (Jensen et al.,
1997; Phillips, Faustman, Lynch, Govoni, Hoagland, &
Zinn, 2001). Jensen et al. (1997) speculated that the
beneficial effect of added vitamin E on muscle colour
depends on the relative muscle vitamin E level in con-
trols. Where these are above a critical level, say 3.5 mg/g,
no beneficial effect is seen. Where the level is low, for
example, in the study of Asghar, Gray, Booren, Gomaa,
Abouzied, and Miller (1991; Table 5), supplementation
to the critical level improves colour retention. In con-
trast, suppression of lipid oxidation is consistently
observed at all muscle levels of added vitamin E.
The total fatty acid content of muscle (i.e. neutral
lipid plus phospholipid fatty acids), termed marbling
fat, has long been recognised as a factor in eating
26 J.D. Wood et al. / Meat Science 66 (2003) 21–32

Table 5 as observed in backfat (Wood, 1984; Wood, Jones,

Effects of supranutritional vitamin E on pork quality (Asghar et al., Bayntun, & Dransfield, 1985). The high percentage of
1991)
18:2 in Durocs was therefore unexpected and they also
Vitamin E in feed (mg/kg) had the highest concentration of 18:3 in neutral lipid.
Values for phospholipid were more similar between the
10 100 200
breeds.
Muscle vitamin E (mg/g) 0.5 2.6 4.7
Drip loss (%)a 20.1 19.5 12.2 3.3. Fatty acids and meat quality in ruminants
Muscle colour (a*)a 7.0 9.3 10.0
TBARS (mg MAD/kg) 3.0 0.94 0.58
Ruminant fat tissue is naturally firmer than that of
a
6 Day display time. pigs, because the fatty acid profile is more saturated. In
the first stages of fattening in cattle, the concentration
quality, especially juiciness and tenderness although the of saturated relative to unsaturated fatty acids increases
strength of the relationship has been questioned (Wood, as in pigs, but beyond a certain fat level in the animal
1990). There are several possible explanations for a this ratio then declines. In very fat cattle, fat is soft and
positive effect of lipid on tenderness, including the oily, mainly due to an increase in 18:1 relative to 18:0
location of neutral lipid in fat cells within the perimy- and 16:0 (Leat, 1975; Wood, 1984). Wood (1984) recor-
sium which could have a physical effect in separating ded a value for 18:0 of 14.7% of total fatty acids in a
muscle fibre bundles, beginning the process of tender- young heifer and 2.7% in an 11-year-old fat steer. In
isation by ‘‘opening up the muscle structure’’. Lipids 1000 lambs selected in four abattoirs and sampled
could also trap moisture in muscle, improving juiciness. throughout the year, Enser and Wood (1993) found that
There are clear genetic effects on the concentrations of the concentration of 18:0, as with pigs, showed the
total fatty acids in muscle, for example, Wood, Whit- highest correlation with melting point (Fig. 2). Average
tington, Nute, Richardson, Sheard, and Chang (in pre- melting point of subcutaneous fat was 39.5  C (range
paration) found that Duroc and Berkshire pure bred 30–49  C) and correlations were 0.89, 0.42 and 0.31
pigs had higher concentrations of both neutral lipids for 18:0, 18:1 and 18:2 fatty acid concentrations,
and phospholipids in longissimus and psoas muscles respectively. It was calculated that only 35% of these fat
than Large Whites and Tamworths (Table 6). The two samples would melt in the mouth of the consumer,
‘‘traditional’’ breeds (Berkshire and Tamworth) had by confirming the hard nature of lamb fat and its tendency
far the fattest carcasses so the results indicated a big to feel sticky in the mouth. The lowest melting points
genetic effect on ‘‘fat partitioning’’ between intramus- were in the summer months May–August corresponding
cular and carcass fat depots. The fatty acid composition with the arrival of ‘‘new seasons’’ lamb.
results were also not as predicted. When neutral lipid In lambs, especially ram lambs, soft fat develops in
increases, the percentage of 18:2 is expected to decline as animals fed grain-based (concentrate) diets. This is due
in the Berkshires compared with the Large Whites and not only to a lower concentration of 18:0 but also to an
increased deposition of medium to long-chain
(C10–C17) branched chain fatty acids formed from
Table 6
Backfat thickness and longissimus neutral lipid and phospholipid fatty methylmalonate, a metabolite of propionate (Busboom
acid content and composition in four pig breeds (Wood et al., in pre- et al., 1981). These authors found that the total con-
paration) centration of branched chain fatty acids was a good
Berkshire Duroc Large White Tamworth Breed effect
predictor of lamb fat firmness but the correlation with
18:0 was equally strong (Table 7). As the concentration
P2 backfat 15.5 9.3 8.0 14.9 *** of branched chain fatty acids increased, that of 18:0 fell,
thickness (mm)
perhaps by direct inhibition.
Neutral lipid The presence of the rumen makes fatty acid composi-
fatty acids tion in beef and sheep more difficult to manipulate by
Weighta 2.29 1.98 0.94 0.91 *** changing diet than in pigs. Nevertheless, there are some
18:2b 6.78 10.14 10.49 8.08 *** clear effects of diet on tissue fatty acid composition.
18:3b 0.58 0.87 0.78 0.67 ***
Scollan, Choi, Kurt, Fisher, Enser, and Wood (2001)
Phospholipid and Vatansever et al. (2000) fed different oils to cattle in
fatty acids a forage:concentrate diet in which total oil content was
Weighta 0.45 0.45 0.39 0.40 *** 6% of dry matter of which 3% was the test oil. Animals
18:2b 29.10 30.10 29.35 31.40 *** were fed for 120 days, after which steaks from long-
18:3b 0.70 0.67 0.61 0.75 ***
issimus and burgers from the forequarter muscles infra-
a
% Of muscle. spinatus, supraspinatus and triceps brachi were examined
b
% Of neutral lipid or phospholipid. (Vatansever et al., 2000). A loin joint was conditioned
 J.D. Wood et al. / Meat Science 66 (2003) 21–32 27

Fig. 2. The relationship between melting point and the concentration of stearic acid in lamb subcutaneous fat (Enser & Wood, 1993).

Table 7 75:25) in sealed plastic containers. Lipid oxidation tests

Correlations between fat firmness, fat colour and fatty acid composi- for the burgers were conducted after heating in a plastic
tion in ram and wether lambs fed high and low energy diets (Busboom bag to 78  C, cooling and 24 h storage. This procedure
et al., 1981)
was used to enhance lipid oxidation and test the stabi-
Fat coloura Fat firmnessb lity of the high-PUFA beef. The results (Fig. 3) show
18:0 0.53 0.81
that n-3 PUFA concentrations were greatly influenced
18:1 0.04 0.12 by diet. The linseed diet doubled the 18:3 concentration
18:2 0.12 0.22 in phospholipids compared with controls, which led to a
BCFAc 0.53 0.81 higher level of EPA but not docasahexanoic acid (DHA
a
1 White to 5 yellow.
22:6; Fig. 3a). These two fatty acids were increased in
b
1 Firm to 5 very soft. the fish oil and linseed/fish oil diets. Lipid oxidation in
c
C10–C17 branched chain fatty acids. the steaks increased at 8 and 11 days of display (Fig. 3b)
and was especially high in the fish oil diet, exceeding the
cut off value of 2 mg malonaldehyde per kg of meat, at
for 11 days at 1  C after which steaks were cut, wrapped which rancidity may be detected by consumers (You-
in oxygen-permeable film and displayed for up to 16 nathan & Watts, 1959). Lipid oxidation in the burgers
days under simulated retail conditions. Lipid oxidation was much more pronounced than in the steaks, a
and colour measurements were made during this time. TBARS value of 2 mg malonaldehyde per kg being
After 5 days of display, steaks were taken for sub- achieved at only 3 days of display (Vatansever et al.,
sequent analysis by the trained taste panel. Burgers 2000). Colour saturation was also affected by feeding
were also used for lipid oxidation and colour measure- treatment, declining fastest in the fish oil diet in which
ments, being packed in a modified atmosphere (O2:CO2 lipid oxidation was greatest (Fig. 3c).
28 J.D. Wood et al. / Meat Science 66 (2003) 21–32

Fig. 3. Effects of dietary n-3 PUFA on fatty acid composition and meat quality in steers. (a) Phospholipid fatty acids in minced forequarter muscles
(%) (Vatansever et al., 2000). (b) Lipid oxidation in longissimus after 11 days conditioning at 1  C and display in overwrapped packs at 4  C.
Assessed as TBARS (mg malonaldehyde/kg) (Vatansever et al., 2000). (c) Colour saturation of burgers prepared from forequarter muscles during
retail display in modified atmosphere packs (Vatansever et al., 2000). (d) Saturated aldehydes (ng/100 g) detected in headspace volatiles of cooked
longissimus (Elmore et al., 1999). (e) Unsaturated aldehydes (ng/100 g) detected in headspace volatiles of cooked longissimus (Elmore et al., 1999). (f)
Taste panel results for grilled longissimus steaks (0–100 scales; Vatansever et al., 2000).
 J.D. Wood et al. / Meat Science 66 (2003) 21–32 29

In order to understand the links between meat com- P:S ratio in meat beyond the 0.1 level normally seen. To
position and flavour, the beef produced from the lin- achieve higher values, two options are to feed a pre-
seed, fish oil and linseed/fish oil diets was analysed for dominantly cereal (concentrate) diet in which rumen
aroma volatile production after cooking in an auto- biohydrogenation is less effective or to ‘‘protect’’ the
clave. The results (Figs. 3d and e) showed that the sam- dietary oil using a procedure such as formaldehyde
ples with high n-3 PUFA concentrations produced treatment of dietary protein which protects the oil
higher concentrations of lipid degradation products, within a matrix structure (Scott, Cook, & Mills, 1971).
particularly saturated and unsaturated aldehydes, alco- We have a fed a 2:1 soya oil:linseed oil supplement in
hols and ketones. Aldehydes were quantitatively the recent work and by so doing have raised the P:S ratio to
most important and since they have low odour thresh- around 3. The n-6:n-3 ratio was raised slightly, but still
olds they are thought to be the reason for the changes in remained below the target value of 4.0 (Enser, Scollan,
flavour of the modified beef (Elmore, Mottram, Enser, Gulati, Richardson, Nute, & Wood, 2001).
& Wood, 1999). Some of these aldehydes are more likely Studies with beef and lamb have shown that the con-
to derive from 18:1 and 18:2 than from 18:3. It is sug- centrations of 18:3 and 20:5 in muscle phospholipids are
gested that free radicals formed from the more unsatu- higher when animals have consumed grass than when
rated and easily oxidised n-3 PUFA initiated the they have been fed grain-based (concentrate) diets. In
oxidation of these more abundant fatty acids (Elmore et the latter case, 18:2 and arachidonic (20:4 n-6) fatty
al., 1999). The taste panel detected a difference in fla- acids are increased relative to 18:3 (Enser et al., 1998;
vour attributes of steaks from the fish oil group and the Fisher et al., 2000; Marmer, Maxwell, & Williams,
rest (Fig. 3f), recording higher values for rancidity and 1984). These results are due to the predominance of 18:3
fishy notes and a lower score for overall liking, however in grass lipids and 18:2 in most other plants and seeds.
these differences were not large on the 100-point scale. In the study of Fisher et al. (2000), Suffolk cross
Campo et al. (2003) have studied the effects of indivi- lambs were reared on lowland grass or on a standard
dual fatty acids on aromas in an in vitro system in concentrate diet. The total lipid of semimembranosus
which the fatty acids were heated alone and with contained higher concentrations of 18:3, 20:5 and 22:6
cysteine and ribose. The results (Table 8) show that in the grass-fed group and higher concentrations of 18:2
meaty aromas are much more pronounced when and 20:4 in the concentrate group (Fig. 4). In grilled loin
cysteine and ribose are also present, i.e. they derive from chops, taste panellists gave higher scores for lamb fla-
interactions between Maillard reaction products and vour and overall liking to the grass group and higher
fatty acids. On the other hand, some flavour notes were scores for abnormal lamb flavour, metallic, bitter and
more associated with the fatty acids alone, e.g. oily. The rancid to the concentrate group.
fatty acids18:1, 18:2 and 18:3 produced different odour In a study involving British lambs fed grass and
profiles, e.g. 18:3 produced high scores for fishy, linseed/ Spanish lambs fed milk and concentrates, analgous dif-
putty and creosote. The term ‘‘grassy’’ was also scored ferences in fatty acid composition were observed, i.e. the
higher for 18:3 plus cysteine and ribose, especially when grass fed animals had higher muscle concentrations of
Fe SO4 was used to catalyse the oxidation reactions. n-3 PUFA and the concentrate-fed animals had higher
Although some dietary PUFA in linseed and other concentrations of the n-6 PUFA (Sanudo et al., 2000).
plant oils escapes rumen biohydrogenation, a high pro- When the meat was assessed by British and Spanish
portion (> 90%) is hydrogenated leading to high values taste panels, both found the British lamb (higher in n-3
for saturated fatty acids in ruminant meats (Scollan, PUFA) to have a higher odour and flavour intensity,
Dhanoa, Choi, Maeng, Enser, & Wood, 2001b). So but whereas the British panel preferred the flavour and
studies such as that by Scollan, Choi et al. (2001a) and overall eating quality of the grass-fed lamb, the Spanish
Vatansever et al. (2000) have usually not increased the panel scored flavour liking and overall liking higher in
the Spanish lamb (Fig. 5). This preference for grain-fin-
Table 8 ished products is also expressed by US taste panellists
Taste panel scores for odours (0–100) in reaction mixtures of fatty and consumers who are more used to the taste of beef
acids, cysteine (c) and ribose (r) (Campo et al., 2003)
and lamb produced in feedlot conditions and prefer it to
Descriptor 18:1 18:2 18:3 18:1 + 18:2 + 18:3 + grass-fed beef and lamb (Kemp, Mahyuddin, Ely, Fox,
c+r c+r c+r & Moody, 1981; Larick & Turner, 1990; Medeiros,
Corned beef 0.6a 0.5a 0.3a 30.3c 18.4b 20.6b Field, Menkhaus, & Russell, 1987). In the British/
Meaty 0.5a 2.9a 0.4a 21.4c 15.7bc 11.0b Spanish data set, wide ranges of n-3 and n-6 PUFA
Oily 18.8b 6.7a 8.7a 3.6a 2.9a 1.4a concentrations were found and quite strong correlations
Fishy 0.1a 0.4a 6.4b 0.1a 0.1a 2.7a were seen between these and the taste panel scores, the
Linseed/putty 1.0a 2.3ab 30.4c 1.6a 2.8b 12.4b
strongest relationships involving 18:2, 20:4 and 18:3
Creasote 0.9a 3.2ab 2.2a 9.7bc 11.1c 17.2c
(Table 9). For example, the concentration of 18:3 was
Means with different letters (a–c) are significantly different (P< 0.05) positively correlated with odour and flavour intensity
30 J.D. Wood et al. / Meat Science 66 (2003) 21–32

Fig. 5. Scores given by British (1–8) and Spanish (0–100) taste panels
to lamb produced on grass in Britain and on concentrates in Spain
Fig. 4. Fatty acid composition and flavour of muscles in sheep fed (Sanudo et al., 2000).
grass or concentrates (Fisher et al., 2000). (a) n-6 and n-3 PUFA as %
total lipid in m. semimembranosus. (b) Taste panel scores for grilled
longissimus steaks (0–100 scales). Table 9
Correlations between fatty acid composition (%) and taste panel
scores given by a Spanish (ESP) and British (GB) taste panel (Sanudo
scores given by both taste panels. The correlations with et al., 2000)
flavour liking and overall liking were positive for the
British panel and negative for the Spanish panel. Flavour intensity Flavour liking
An explanation for these positive relationships ESP GB ESP GB
between 18:3 and meat flavour is that the oxidation
18:2 n-6 0.5 0.6 0.7 0.5
products of 18:3 and it’s derivatives are directly respon- 18:3 n-3 0.4 0.7 0.7 0.6
sible for the differences in flavour observed between
grass-fed cattle and sheep. Larick and Turner (1990)
also found that the headspace volatile compounds pro- compared with a grain diet. Grass feeding also increased
duced on cooking beef fed grass or grains were domi- concentrations of diterpenoids which derive from
nated by lipid oxidation products, especially aldehydes chlorophyll breakdown in the rumen. Similar results
and Priolo, Mikol, and Agabaiel (2001) concluded in a have been found in cattle (Melton, 1990). Young et al.
review that n-3 PUFA oxidation products are mainly (1997) also found that the concentration of 3-methyl-
responsible for the particular flavour of grass-fed lamb. indole (skatole), which is responsible for boar taint in
However, another explanation for the results is that n-6 pigs, was increased in grass diets and was partly
and n-3 PUFA are markers for grain and grass diets, responsible for the grass-fed effect.
respectively and other components of meat are more Several authors have shown that lipid oxidation and
directly responsible for the flavours produced. In sheep, colour development in ruminant meats is influenced by
several authors have shown that branched chain fatty both fatty acid composition and the concentration of
acids of medium chain length are important constituents the tissue antioxidant, vitamin E (Arnold, Arp, Scheller,
of lamb odours and flavours (Wong, Nixon, & Johnson, Williams, & Schaefer, 1993; Renerre, 2000). In a recent
1975) and Young, Berdagué, Viallon, Rousset-Akrim, comparison of grass-fed and grain-fed cattle by our
and Theriez (1997) showed that 4-methyloctanoic acid group (Warren et al., 2002), the bright red colour asso-
and 4-methylnonanoic acid were increased on a grass ciated with oxymyoglobin was retained longer in retail
 J.D. Wood et al. / Meat Science 66 (2003) 21–32
display in the grass-fed group. Although the total con-
centration of unsaturated fatty acids was similar in
both, grass feeding produced higher concentrations of
more oxidisable n-3 PUFA and grain feeding increased
levels of n-6 PUFA. The conclusion was that anti-
oxidants in grass probably caused higher tissue levels of
vitamin E in these animals with benefits for lower lipid
oxidation and better colour retention despite greater
potential for lipid oxidation. Yang, Brewster, Lanari,
and Tume (2002) reported high tissue levels of vitamin
E in grass-fed beef, similar to those recorded in grain-
fed cattle supplemented with 2.5 g vitamin E/head/day.
In studies with sheep, Kasapidou, Wood, Sinclair,
Wilkinson, and Enser (2001) showed that low tissue
concentrations of vitamin E are associated with lower
amounts of both n-6 and n-3 PUFA in tissues. This
suggests that loss of PUFA occurs in vivo when anti-
oxidant status is low. These studies also showed that
muscle concentrations of vitamin E in sheep were often
well below the level of 3.0–3.5 mg/g suggested by Arnold
et al. (1993) and Liu, Scheller, Arp, Schaefer, and Wil-
liams (1996) as necessary for optimum antioxidant sta-
tus in cattle. This was particularly true when the lambs
were fed a grain-based (concentrate) diet alone. When
grass was included in a mixed diet with concentrates,
vitamin E levels were usually restored to around 3 mg/g.
The associations found between lipid oxidation, colour
development and vitamin E concentration in steers by
Liu et al. (1996) show that the interrelationship between
fatty acid composition, in both lipid fractions and tissue
vitamin E in ruminants is crucial to many aspects of
metabolism, with eventual implications for meat quality.
Acknowledgements
We are grateful to our collaborators in providing
many of the results presented here, including Institute of
Grassland and Environmental Research, Harper Adams
University College and University of Reading. Funding
was provided by Department for Environment, Food
and Rural Affairs (DEFRA), Meat and Livestock
Commission, ABN Ltd, JSR Farms Ltd, Tesco Stores
Ltd, Roche Products Ltd, International Fishmeal and
Oil Manufacturers Association and Southern Counties
Fresh Foods Ltd. We gratefully acknowledge the tech-
nical assistance of Kathy Hallett, Fran Whittington,
Kevin Gibson, Anita Robinson, Rose Ball, Anne Baker
and Sue Hughes
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